A Taxonomic Revision of Bromus (Poaceae: Pooideae: Bromeae) in México and Central America

Home , Arbutus tessellata, Bromus, Bromus anomalus, Bromus arizonicus, Bromus berteroanus, Bromus carinatus

http://dx.doi.org/10.11646/phytotaxa.185.1.1 PHYTOTAXA

185

A taxonomic revision of Bromus (Poaceae: Pooideae: Bromeae) in México and Central America

JEFFERY M. SAARELA1,4, PAUL M. PETERSON2 & JESUS VALDÉS-REYNA3 1Botany Section, Research & Collections Services, Canadian Museum of Nature, PO Box 3443, Station D, Ottawa, Ontario K1P 6P4, Canada; [email protected] 2Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C., 20013-7012, U.S.A.; [email protected] 3Departamento de Botanica, Univerisidad Autonoma Agraria "Antonio Narro" Buenavista, Saltillo, Coahuila 25315, MÉXICO; jvaldé[email protected] 4Author for correspondence

Magnolia Press Auckland, New Zealand

Accepted by MARIA VORONTSOVA: Jun. 2014; published: 14 Nov. 2014 1 JEFFERY M. SAARELA, PAUL M. PETERSON & JESUS VALDÉS-REYNA A taxonomic revision of Bromus (Poaceae: Pooideae: Bromeae) in México and Central America (Phytotaxa 185) 147 pp.; 30 cm. 14 November 2014 ISBN 978-1-77557-577-1 (paperback) ISBN 978-1-77557-578-8 (Online edition)

FIRST PUBLISHED IN 2014 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/phytotaxa/

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Abstract ...... 3 Resumen...... 4 Introduction...... 4 Taxonomic History of Bromus in México and Central America ...... 4 Materials and Methods...... 6 Taxonomic Treatment ...... 7 Bromus Linnaeus (1753: 76), nom. et. typ. cons...... 7 A new Bromus section endemic to México...... 7 Key to Bromus of México and Central America...... 8 Clave para las especies de Bromus en México y América Central...... 9 1. Bromus anomalus Ruprecht ex Fournier (1886: 126). Figs. 1–3...... 10 2. Bromus arizonicus (Shear) Stebbins in Stebbins et al. (1944: 309). Figs. 5, 6...... 24 3. Bromus attenuatus Swallen (1950: 397). Figs. 8, 9...... 28 4. Bromus berteroanus Colla (1836: 25, pl. 58). Figs. 11, 12...... 31 5. Bromus carinatus Hooker & Walker Arnott (1840: 403)...... 35 5a. Bromus carinatus var. carinatus. Figs. 15A–D, 16...... 42 5b. Bromus carinatus var. marginatus (Nees) Barkworth & Anderton in Barkworth et al. (2006: 240). Figs. 15E–G, 18, 19... 47 6. Bromus catharticus Vahl (1791: 22). Figs. 21–23...... 61 7. Bromus densus Swallen (1950: 396). Figs. 25–27...... 68 8. Bromus diandrus Roth (1787: 44). Figs. 29, 30...... 72 9. Bromus dolichocarpus Wagnon (1950: 65). Figs. 32, 33...... 77 10. Bromus exaltatus Bernhardi (1841: 90). Figs. 35, 36...... 83 11. Bromus frondosus (Shear) Wooton & Standley (1912: 144). Figs. 38, 39...... 88 12. Bromus hordeaceus Linnaeus (1753: 77). Figs. 41, 42...... 92 13. Bromus inermis Leysser (1761: 16). Figs. 44, 45...... 96 14. Bromus japonicus Houttuyn (1772: 315). Figs. 47, 48...... 98 15. Bromus lanatipes (Shear) Rydberg (1906: 52). Fig. 50, 51...... 101 16. Bromus madritensis Linnaeus (1755: 5). Figs. 53, 54...... 104 17. Bromus pinetorum Swallen (1943: 77). Fig. 56...... 108 18. Bromus pseudolaevipes Wagnon (1950: 64). Figs. 58, 59...... 110 19. Bromus richardsonii Link (1833: 281). Figs. 61, 62...... 113 20. Bromus rubens Linnaeus (1755: 5). Figs. 64, 65...... 122 21. Bromus secalinus Linnaeus (1753: 76). Fig. 67...... 128 22. Bromus tectorum Linnaeus (1753: 77). Figs. 69, 70...... 129 Excluded Taxa...... 133 Acknowledgements...... 134 References...... 134

Abstract

A taxonomic treatment of Bromus in México and Central America is presented. Twenty-two species and one variety are recognized in the region. Twelve of these species are native, and ten are adventive and introduced. Bromus section Mexibromus is newly described, representing a distinct lineage in molecular studies of the genus. It includes two species endemic to México (B. attenuatus, B. densus) and one endemic to México and Guatemala (B. dolichocarpus), distinguished from species in other Bromus sections by their 3(–5)-nerved lemmas. Bromus pinetorum is a poorly understood species of Bromus sect. Bromopsis endemic to Coahuila. Taxa that have been variously treated in the past are discussed in detail. Bromus mucroglumis and B. thysanoglottis are treated as synonyms of B. richardsonii, and B. meyeri is treated as a synonym of B. anomalus. Two varieties are recognized in B. carinatus, B. carinatus var. carinatus and B. carinatus var. marginatus. Lectotypes are designated for the names B. carinatus var. californicus, B. hookeri var. schlechtendalii, B. trinii, B. trinii var. pallidiflorus and B. trinii var. stricta. A second-step neotype is designated for B. rigidus. Keys for determining the species in English and Spanish, descriptions, synonymies, illustrations, distribution maps, comments and lists of all specimens examined are provided for each species.

A TAXONOMIC REVISION OF BROMUS (POACEAE) Phytotaxa 185 (1) © 2014 Magnolia Press • 3 collector given on the specimen label) and originally determined as B. commutatus; the specimen is B. japonicus. This species was again reported recently from the same experimental station by Royo Márquez & Melgoza Castillo (2012), likely based on the same mis-identified specimen.

Bromus grandis (Shear 1900: 243) Hitchcock in Jepson (1912: 175) Wiggins (1980) reported this taxon from N Sierra Juárez, Baja California. Gould & Moran (1981) noted that they did not see specimens of this taxon and did not treat it. We have not seen specimens of this taxon from Baja California or elsewhere in México.

Bromus orcuttianus Vasey (1885a: 223) Wiggins (1980: 912) reported this taxon from "northernmost Baja California". Gould & Moran (1981) noted that they did not see specimens of this taxon and did not treat it. We have not seen specimens of this taxon from Baja California or elsewhere in México. A specimen determined originally as B. orcuttianus from southwestern Chihuahua (Aug–November 1883, Palmer 233, NY!) is a species of Brachypodium. Another specimen from Chihuahua (Jones s.n., RSA-POM), incorrectly determined as B. orcuttianus, is B. carinatus var. marginatus.

Bromus porteri (Coulter 1885: 425) Nash (1895: 512) See comments under B. anomalus.

Bromus sterilis Linnaeus (1753: 77) Espejo-Serna et al. (2000) reported B. sterilis from Baja California. This report may be based on Paulin & Morales 218 (MEXU-1098059) and MEXU-109768, both collected in Baja California and originally determined as B. sterilis; the former specimen is B. carinatus, the latter is B. madritensis.

Bromus texensis (Shear 1900: 41) Hitchcock (1913: 381) Reported from México by Hitchcock (1913), on the basis of a specimen in US (Pringle 2052). Wagnon (1952) noted that this specimen is B. anomalus, a determination with which we agree.

Acknowledgements

We thank the National Geographic Society Committee for Research and Exploration (Grant No. 8848-10, 8087- 06), the Smithsonian Institution’s Restricted Endowments Fund, the Scholarly Studies Program, Research Opportunities, Atherton Seidell Foundation, Biodiversity Surveys and Inventories Program and Small Grants Program, and the Canadian Museum of Nature, all for financial support. Micheline Bouchard and Jennifer Doubt (CAN) facilitated loans, and we are grateful to the curators of the herbaria from which loans were obtained, and to Laura Crumbacher who provided digital images from ARIZ. Paul Sokoloff helped prepare the illustrations for publication. Rich Rabeler (MICH), Alan Prather (MSC) and Jon Rebman (SD) provided permission to publish images of specimens from their respective herbaria. Richard Felger provided information on Bromus arizonicus. We are grateful to Yolanda Herrera Arrieta and M. Soccoro González-Elizondo for their generous hospitality during our numerous field trips to México. We would also like to acknowledge two anonymous reviewers for suggesting improvements to the manuscript.

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